Overview

Distribution

Range Description

This species occurs in Bangladesh, Brunei, Cambodia, India (Nicobar Islands), Indonesia (including Bali, Bangka, the Batu Islands, Bawean, Belitung, Java, Kalimantan Borneo, the Kangean Islands, Karimata, Karimunjawa, Lingga, Lombok, the Natuna Islands, Nias, Nusatenggara, the Riau Archipelago, Simeulue, Sumatra, Sumba, Sumbawa, and Timor), Lao PDR, Malaysia (including the Peninsula as well as Sabah and Sarawak Borneo), Myanmar (including the Mergui Archipelago), the Philippines (Balabec, Basilan, Cagayan Sulu, Culion, Jolo, Leyte, Luzon, Mindanao, Mindoro, Palawan, and Samar), Singapore, Thailand (including offshore islands), Timor-Leste and Viet Nam. On mainland Southeast Asia, there is a hybrid zone with Macaca mulata in central areas that makes it difficult to determine the northern limits of its range. The species has been introduced to Kabeana Island, Indonesia, the Pacific island of Palau, Mauritius in the Indian Ocean, and to New Guinea (Groves 2001) (these introduced populations are not included in the distribution map).

M. f. fascicularis
Occurs in Brunei, Cambodia, Indonesia (including Bali, Bangka, the Batu Islands, Bawean, Java, Kalimantan Borneo, Karimata, Lingga, the Natuna Islands, Nias, Nusatenggara, the Riau Archipelago, Sumatra, Sumba, and Timor), Malaysia (including the peninsula as well as Sabah and Sarawak Borneo), the Philippines (western Mindanao and the Sulu Archipelago), Singapore, southern Thailand (including offshore islands), and southern Viet Nam (Groves 2001).

M. f. aureus
Occurs in southernmost Bangladesh, Lao PDR, Myanmar (including the Mergui Archipelago) and west-central Thailand (Groves 2001).

M. f. umbrosus
Occurs on Nicobar Islands of India (Little Nicobar, Great Nicobar and Katchall), where it is found up to 600 m (Umapathy et al. 2003; Molur et al. 2003; Groves 2001).

M. f. fuscus
Occurs on Pulau Simeulue, off the northwestern coast of Sumatra, Indonesia (Groves 2001).

M. f. condorensis
Occurs on Con Son Island, off the coast of southern Viet Nam (Groves 2001).

M. f. tua
Occurs on Pulau Maratua, east of Borneo, Indonesia (Groves 2001).

M. f. philippensis
Occurs in the Philippines. Found on the islands of Balabac, Basilan, Biliran, Bohol, Busuanga, Camiguin, Catanduanes, Culion, Leyte, Luzon, northeastern Mindanao, Mindoro, Negros, Panay, Palawan, Samar and Sibuyan (Groves 2001; Heaney et al. 1998).

M. f. lasiae
Occurs on Pulau Lasia, off the northewestern coast of Sumatra, Indonesia (Groves 2001).

M. f. karimondjawae
Occurs on Pulau Karimunjawa and (probably) nearby Pulau Kemujan, Java Sea, Indonesia (Groves 2001).

M. f. atriceps
Occurs on Khram Yai Island, off the southeastern coast of Thailand (Groves 2001).
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Geographic Range

Long tailed or crab eating macaques (Macaca fascicularis) are found in southeast Asia from Burma to the Philippines and southward through Indochina, Malaysia, and Indonesia. They are found as far east as the Timor Islands.

Biogeographic Regions: oriental (Native )

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Physical Description

Morphology

Physical Description

The body fur of long-tailed macaques tends to be grey-brown to reddish brown. These colors are always paler ventrally. The face is brownish-grey with cheek whiskers. The eyes are directed forward for binocular vision. The nose is flat and the nostrils are narrow and close together (catarrhine condition). Long-tailed macaques have shovel-shaped incisors, conspicuous canines, and bilophodont molars. The tooth formula is I 2/2, C 1/1, PM 2/2, and M 3/3.

The body length, not including the tail, is 40 to 47 cm. The greyish- brown or reddish colored tail is 50 to 60 cm. Long-tailed macaques exhibit sexual dimorphism in size. The average weight for males is 4.8 to 7 kg and 3 to 4 kg for females, approximately 69% of average male weight.

Range mass: 3 to 7 kg.

Range length: 40 to 47 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
The species is extremely tolerant of a range of habitats, including mangrove and swamp forests, and can be found in agricultural areas near forest (secondary growth, secondary forest, and primary forest) (Thomas 1898; Fooden 1991, 1995; Rabor 1986; Goodman and Ingle 1993; Heaney et al. 1991; Rickart et al. 1993; Danielsen et al. 1994). On the Nicobars, M. f. umbrosus is found in mangroves and coastal forests dominated by Pandanus species (Molur et al. 2003). M. f. aureus is found in evergreen forests and coastal mangroves (Molur et al. 2003). The species has been reported as occurring to elevations of up to 1,000 m on Java, Borneo, and Sumatra (Indonesia), and up to 1,800 m in the Philippines (Heaney et al. 1998). On the mainland, it generally occurs at lower elevations: up to 700 m in Thailand, while in Cambodia and Viet Nam it generally occurs below 300 m. This species is semi-terrestrial, diurnal, and omnivorous (Molur et al. 2003).

Systems
  • Terrestrial
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Long-tailed macaques are "ecologically diverse." Some of the habitats in which they have been found are primary forests, disturbed and secondary forests, and riverine and coastal forests of nipa palm and mangrove. Long-tailed macaques live most successfully in disturbed habitats and on the periphery of forests.

In Sumatra, they achieve their highest population densities in mixed mangrove swamps, secondary hill forests, and riverine forests. Some were also observed in freshwater swamps, scrub grassland, lowland primary forests, and rubber groves.

In Thailand, long-tailed macaques occur in evergreen forests, bamboo forests, and in deciduous forests.

In Malaysia, they are abundant in coastal lowland forests.

This species has been observed drinking much water and eating crabs, they often live near bodies of water. Of the various habitats occupied by long-tailed macaques, the swamp forests seem to have the highest density of them.

In recent years, habitat alteration has expanded the range of some populations of long-tailed macaques. In Malaysia, cleared land, such as plantation areas, has been colonized by this species. It has been observed that some disturbed habitats have higher troop and population sizes than some pristine forests.

This species has the highest degree of arboreality of all macaque species. One study of long-tailed macaque behavior reported that they never came to the ground except within 5 m of the edge of a river near their tree. The population densities of this diurnal species vary from 10 to 400 per squared kilometer.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest ; scrub forest

Aquatic Biomes: coastal ; brackish water

Wetlands: swamp

Other Habitat Features: agricultural ; riparian

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Trophic Strategy

Food Habits

Long-tailed macaques are omnivores, and exploit many different food types, reflecting the diversity of habitats they can utilize. The average length of feeding bouts is 18.3 minutes. There may be on average of twenty bouts per day. They eat a wide variety of foods such as fruits, crabs, flowers, insects, leaves, fungi, grasses, and clay. Clay may be eaten for the potassium found in it, although the potassium levels in the clay are low. However, 96% of the feeding time per day is spent eating fruit. Some limited observations suggest that long-tailed macaques select fruit based on ripeness, which is based on color.

Animal Foods: insects; aquatic crustaceans

Plant Foods: leaves; fruit; flowers

Other Foods: fungus

Primary Diet: omnivore

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Associations

Ecosystem Roles

To the extent that these animals are prey for other species, they may affect populations of those species. Macaca fascicularis may be an important predator in its ecosystem, and may have some impact on prey species.

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Predation

The predators of this species have not been reported.

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Life History and Behavior

Behavior

Communication and Perception

As in other species of the genus Macaca, it is likely that these animals use a combination of visual (facial expression, body postures), auditory (vocalizations), physical (grooming, play, mating, aggression) and possible chemical (olfactory) signals as a part of their repertoire of communication.

Communication Channels: visual ; tactile ; acoustic ; chemical

Perception Channels: visual ; acoustic

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Life Expectancy

Lifespan/Longevity

Although the lifespan of this species has not been reported, it is likely to resemble that of other members of the genus, which seem able to obtain an maximum age of around 30 years in captivity.

Average lifespan

Sex: female

Status: captivity:
25.0 years.

Average lifespan

Status: captivity:
38.0 years.

Average lifespan

Status: captivity:
37.1 years.

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Lifespan, longevity, and ageing

Maximum longevity: 39 years (captivity) Observations: One wild born male lived 37.1 years in captivity, making it as old as 39 when he died (Richard Weigl 2005).
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Reproduction

In Sumatra, each social group of these monkeys contains an average of 5.7 males and 9.9 adult females. Although females outnumber males, indicating that males have more mates available to them than do females, both sexes may mate with multiple partners. However, the alpha male, or highest ranking male, of the group has the greatest access to mates of any males, and probably sires the most offspring.

Mating System: polygynous ; polygynandrous (promiscuous)

Males reach sexual maturity at approximately 6 years of age, whereas females mature by about 4 years of age. Higher-ranking females become reproductively mature before lower-ranking females. The offspring of these higher-ranking females have a greater chance of survival than do the offspring of lower-ranking females. Both of these phenomena are related to the greater food availability and decreased amount of aggression experienced by the higher ranking females.

Female long-tailed macaques show a conspicuous cyclicity of sexual behavior during their menstrual cycle. As they approach ovulation, females experience a swelling of the skin in the perineal region. However, there has been no direct correlation between the frequency of copulation and the degree of swelling of genital tissues. This concealment of ovulation could exist in order to persuade consorting males to stay with females longer.

As a side effect of the prolonged estrus period and concealment of ovulation, it is difficult for humans to estimate the length of gestation. However, it seems likely that the gestation averages approximately 162 days. The average interbirth intervasl is approximately 390 days, indicating that females can produce young each year if high ranking, and every other year otherwise.

Young are nursed until they are approximately 420 days of age. The average interbirth interval is reported to be 390 days, which is less than the average time until weaning. Both the length of nursing and the interbirth interval may be affected by the rank of the mother.

It is known that the births in this species peak from May to July, corresponding with the rainy season.

Breeding interval: Females have an average interbirth interval of 390 days.

Average number of offspring: 1.

Average gestation period: 162 days.

Average weaning age: 420 days.

Average age at sexual or reproductive maturity (female): 4 years.

Average age at sexual or reproductive maturity (male): 6 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous

Average birth mass: 320 g.

Average gestation period: 165 days.

Average number of offspring: 1.

Average age at sexual or reproductive maturity (male)

Sex: male:
1544 days.

Average age at sexual or reproductive maturity (female)

Sex: female:
1238 days.

As in all primates, there is an extended period of juvenile dependence. Females provide the bulk of the care for offspring. The young receive nourishment, encouragement, grooming, protection, and teaching from the mother. Young females are likely to remain in the troup into which they were born, whereas males are likely to emmigrate at or near sexual maturity.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); post-independence association with parents; extended period of juvenile learning; inherits maternal/paternal territory; maternal position in the dominance hierarchy affects status of young

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Macaca fascicularis

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 2 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

GATCGTTGACTCTTTTCAACAAACCATAAGGACATCGGAACCCTGTATTTACTATTTGGTGCATGAGCTGGGGTCATAGGCACTGCCCTG---AGCCTCCTCATTCGAGCTGAACTAGGCCAACCCGGCAGCCTACTAGGCAAC---GACCACATCTACAACGTCATTGTAACGGCCCATGCATTCGTTATAATTTTCTTTACAGTTATACCCATTATAATCGGAGGGTTCGGGAACTGACTAGTGCCCCTAATA---ATCGGCGCTCCCGACATAGCATTTCCCCGTCTAAACAATATAAGCTTTTGACTCCTCCCCCCTTCTTTCCTGCTACTAATTGCATCAGCCGTAGTAGAAGCTGGTGCCGGAACAGGCTGAACAGTATATCCCCCTCTAGCAGGAAACTTCTCCCACCCAGGAGCTTCTGTAGATTTGATT---ATCTTCTCCCTCCACTTAGCAGGTATTTCCTCCATCTTAGGAGCCATCAACTTTATTACCACTATTATCAACATAAAACCTCCCGCAATATCCCAATACCAGACTCCTTTATTTGTCTGATCAATCTTAATCACGGCTATCCTCCTACTACTCTCTCTACCAGTCCTAGCCGCC---GGCATTACCATGTTACTAACAGATCGCAACCTCAACACTACTTTCTTTGATCCTGTTGGAGGAGGAGATCCTATCTTATACCAACACCTATTTTGATTCTTCGGTCACCCCGAAGTCTACATCCTTATTCTTCCCGGCTTTGGGATAGTCTCCCACATTGTAACCTACTACTCTGGAAAAAAA---GAACCATTTGGGTACATGGGTATAGTTTGAGCCATAATATCAATTGGGTTTTTAGGTTTCATTGTATGAGCCCACCACATATTTACAGTTGGCATAGACGTAGATACACGAGCCTATTTCACTTCCGCCACTATAATCATTGCAATCCCCACCGGTGTGAAAGTTTTTAGCTGGCTC---GCTACACTTCATGGAGGC---AACATTAAATGATCCCCAGCAATACTTTGAGCCCTGGGCTTTATCTTCCTATTTACCGTGGGAGGTTTAACCGGCATTATCTTGGCAAACTCATCCCTAGATATCGTACTACACGACACATACTACGTTGTCGCCCACTTCCATTATGTT---TTATCAATAGGGGCTGTATTTGCCATTATAGGAGGCTTTATACATTGATTTCCTTTATTCTCAGGCTATACACTAAACCAAACTTGTGCCAAAGCCCACTTTATCATCATATTCGTGGGCGTAAATTTAACTTTCTTTCCACAACACTTCCTTGGCCTGTCCGGAATACCCCGA---CGCTATTCTGATTATCCTGATGCCTACACC---ACATGAAACATCCTATCATCTATAGGCTCCTTCATCTCACTAGTAGCAGTAATCTTAATAATCTACATGATCTGAGAAGCCTTTGCTTTAAAACGTAAAGTA---CTACTAATCGAACAACCCCTTACTAGT
-- end --

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Statistics of barcoding coverage: Macaca fascicularis

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 2
Specimens with Barcodes: 4
Species With Barcodes: 1
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Ong, P. & Richardson, M.

Reviewer/s
Mittermeier, R.A. & Rylands, A.B. (Primate Red List Authority)

Contributor/s

Justification
This species is listed as Least Concern in view of its wide distribution, presumed large population, tolerance of a broad range of habitats, occurrence in a number of protected areas, and because it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category. Although it is under heavy hunting pressure for meat, sport and trophies, this is not considered a major threat to the species overall.
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There are nine national parks, nine reserves, and two sanctuaries in which some long-tailed macaques reside. Regardless of the type of habitat, there must be at least 500 squared kilometers of habitat necessary to support a viable population of 5,000 long-tailed macaques. This is the minimum size for a reserve for this species.

Long-tailed macaques receive some protection in temple ruins in Thailand and protection and food in temples in Bali. In Malaysia, long-tailed macaques are legally protected, and they are fed and protected in urban forests and parks. In the Philippines, there is much interest in protecting this species. In Indonesia, the species is well-protected, but some of the reserves are being considered for oil drilling and harvesting. Some people in Bali, in fact, consider these primates to be sacred. This may increase the chances of their survival in these reserves. In Thailand, long-tailed macaques may be hunted, captured, or kept in captivity only under license. The export of this species is regulated by a quota system.

Although there are some sanctuaries for long-tailed macaques, hunting is still a problem. In Thailand and Borneo, they are hunted for food. This species is also killed because it is a pest to agriculture. The fact that these macaques destroy crops has prevented some governments from making serious conservation efforts. Long-tailed macaques are collected for medical research. They are one of the five most used primate species in medical research. Many of these macaques were exported to the United States and Great Britian. Habitat loss in these organisms is occurring due to extensive logging operations.

US Federal List: no special status

CITES: appendix ii

IUCN Red List of Threatened Species: least concern

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Population

Population
This is a widespread and often abundant species, and is sometimes commensal with humans. In the Philippines, it ranges from being locally common to uncommon, though this is largely dependent upon hunting pressure. In Bangladesh, the population has been estimated at less than 100 total individuals, and a population on the Teknaf Peninsula has been completely decimated due to development activities (ship-building) (Molur et al. 2003).
There is a paucity of information available on the population status of several island forms. Molur et al. (2003) estimated about 4,800 individuals of M. f. umbrosus, but nothing is known about the status following the December 2004 tsunami. The islands of Katchall and Little Nicobar were submerged during the tsunami, and it is believed to have affected the habitat of this taxon; on the other hand, Shankaran et al. (2005) state that these macaques were not affected seriously on Greater Nicobar Island. The disaster could have impacted their populations on these two islands due to the fruiting Pandanus being affected by the tsunami (M. Singh pers. comm.). The population on Con Son (M. f. conderensis) likely numbers less than 1,000 individuals, and a number of other island forms probably have a similar status.

Population Trend
Decreasing
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Threats

Major Threats
Across much of the range, the major threat to the species is hunting. In mainland Southeast Asia, such as in Cambodia and Viet Nam, females are taken into breeding facilities and males are exported internationally primarily for use in laboratory research. A potential related issue is the release of confiscated long-tailed macaques from the border area of Viet Nam and China (which is where most of the current international trade is being recorded) into the native range of other macaque species. In the Philippines, the species is subject to a high level of hunting, where it is taken for local consumption and hunted for sport. It is also persecuted as a pest. Habitat loss is also a localised threat, but the species can persist in a variety of habitats and very adaptable.
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Management

Conservation Actions

Conservation Actions
The species is included in Appendix II of CITES. It is listed on Schedule I, Part I, Indian Wildlife (Protection) Act, 1972 amended up to 2002, and Schedule III, Bangladesh Wildlife (Protection) Act A 1974 (Molur et al. 2003). In Myanmar it is a “normally” protected species. It is a protected on Appendix 2B on Decree 32 (2006) in Viet Nam. It occurs in many protected areas throughout its range and is relatively common in captivity (M. Richardson pers. comm.).

There is a need for further survey work to assess the current population status of the various island forms. In particular, following the Indian Ocean Island tsunami of December 2004, a targeted survey is required to establish their current status on the Nicobars.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Long-tailed macaques have been known to feed in cultivated fields on such items as young dry rice, cassava leaves, rubber fruit, taro plants, and other crops. They also take food from graveyards, garbage cans, and garbage pits. They have also become involved in aggresive interactions with people.

Negative Impacts: injures humans (carries human disease); crop pest

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Economic Importance for Humans: Positive

Long-tailed macaques, along with other species of macaques, have benefited humans through their use as research models in immunology, surgery, toxicology, and pharmacology. They are also important members of ecosystems and may serve as a basis for ecotourism ventures. They are sometimes still hunted for food.

Positive Impacts: food ; ecotourism ; research and education

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Wikipedia

Crab-eating macaque

The crab-eating macaque (Macaca fascicularis), also known as the long-tailed macaque, is a cercopithecine primate native to Southeast Asia. It is referred to as the cynomolgus monkey in laboratories.[2] It has a long history alongside humans;[7] they have been alternately seen as agricultural pests,[8] sacred animals in some temples,[9] and more recently, the subject of medical experiments.[7] The crab-eating macaque lives in matrilineal social groups with a female dominance hierarchy,[10] and male members leave the group when they reach puberty.[11] They are opportunistic omnivores [12] and have been documented using tools to obtain food in Thailand and Myanmar.[13] The crab-eating macaque is a known invasive species and a threat to biodiversity in several locations including Hong Kong and western New Guinea.[2] The significant overlap in macaque and human living space has resulted in greater habitat loss,[7] synanthropic living, and inter- and intra-species conflicts over resources.

Etymology[edit]

The scientific name of the crab-eating macaque is Macaca fascicularis. Macaca comes from the Portuguese word macaco, which was derived from makaku, a Fiot (West African language) word (kaku means 'monkey' in Fiot).[14] Fascicularis is Latin for 'a small band or stripe'. Sir Thomas Raffles, who gave the animal its scientific name in 1821, did not specify what he meant by the use of this word.

In Indonesia and Malaysia, M. fascicularis and other macaque species are known generically as kera, possibly because of the high-pitched cry of the animal.[15]

The crab-eating macaque has several common names. It is often referred to as the 'long-tailed macaque' due to its tail which is often longer than its body.[16] The name 'crab-eating macaque' refers to its being often seen foraging beaches for crabs. Another common name for M. fascicularis is the 'cynomolgus monkey', which literally means "dog-skin' or 'dog-hide' monkey; this name is commonly used in laboratory settings.

Taxonomy[edit]

There are 10 subspecies of Macaca fasciularis:

Physical characteristics[edit]

The body length of the adult, which varies among subspecies, is 38–55 cm (15–22 in) with relatively short arms and legs. Males are considerably larger than females, weighing 5-9 kilograms (11-20 lb) compared to the 3–6 kg (7-13 lb) of females.[16] The tail is longer than the body, typically 40–65 cm (16–26 in), which is used for balance when they jump distances up to 5 meters (16.4 feet).[16] The upper parts of the body are dark brown with light golden brown tips. The under parts are light grey with a dark grey/brown tail. Crab-eating macaques have backwards directed crown hairs which sometimes form short crests on the mid-line. Their skin is black on their feet and ears, whereas the skin on the muzzle is a light grayish pink color. The eyelids often have prominent white markings and sometimes there are white spots on the ears. Males have a characteristic mustache and cheek whiskers, while females have only cheek whiskers. Crab-eating macaques have a cheek pouch which they use to store food while foraging. Females show no perineal swelling.[18]

Behavior[edit]

Group living[edit]

Batu Caves, Malaysia

Macaques live in social groups that contain three to twenty females, their offspring, and one or many males. The groups usually have fewer males than females. In social groups of macaques, there is a clear dominance hierarchy among females. These ranks remain stable throughout the female’s lifetime and also can be sustained through generations of matrilines. Females have their highest birth rates at approximately ten years of age and completely stop bearing young by age twenty-four.[10]

The social groups of macaques are female-bonded, meaning that males will disperse at the time of puberty. Thus, group relatedness on average appears to be lower than compared to matrilinearity | matrilines. There is more difference in relatedness when comparing high-ranking lineages to lower ranking lineages, with higher-ranking individuals being more closely related to one another. Additionally, groups of dispersing males born into the same social groups display a range of relatedness, at times appearing to be brothers while at other times appearing to be unrelated.[11]

In addition to the matrilineal dominance hierarchy, there are also male dominance rankings. Alpha-males have a higher frequency of mating compared to their lower-ranking conspecifics. The increased success is due partially to his increased access to females and also due to female preference of an alpha male during periods of maximum fertility. Though females have a preference for alpha males, they do display promiscuous behavior. Through this behavior, females risk helping to rear a non-alpha offspring, yet benefit in two specific ways, both in regard to aggressive behavior. First, there is decreased value placed on one single copulation. Moreover, the risk of infanticide is decreased due to the uncertainty of paternity.[19]

Increasing group size leads to increased competition and energy spent trying to forage for resources, and in particular, food. Further, social tensions build and the prevalence of tension-reducing interactions like social grooming fall with larger groups. Thus, group living appears to be maintained solely due to the safety against predation.[20]

Conflict[edit]

Group living in all species is dependent on tolerance of other group members. In crab-eating macaques, successful social group living maintains that there must be post-conflict resolution. Usually, less dominant individuals will lose to a higher-ranking individual when conflict arises. After the conflict has taken place, lower-ranking individuals tend to fear the winner of the conflict to a greater degree. In one study, this was seen by the ability to drink water together. Post-conflict observations showed a staggered time between when the dominant individual begins to drink and the subordinate. Long-term studies reveal that the gap in drinking time closes as the conflict moves further into the past.[21]

Grooming and support in conflict among primates is considered to be an act of reciprocal altruism. In crab-eating macaques, an experiment was performed in which individuals were given the opportunity to groom one another under three conditions: after being groomed by the other, after grooming the other, and without prior grooming. After a grooming took place, the individual who received the grooming was much more likely to support its groomer than one who had not previously groomed that individual. These results support the reciprocal altruism theory of grooming in long-tailed macaques.[22]

Crab-eating macaques demonstrate two of the three forms of suggested post-conflict behavior. In both captive and wild studies, the monkeys demonstrated reconciliation, or an affiliative interaction between former opponents, and redirection, or acting aggressively towards a third individual. Consolation was not seen in any study performed.[23]

Post-conflict anxiety has been reported in crab-eating macaques that have acted as the aggressor. After a conflict within a group, the aggressor appears to scratch himself at a higher rate than before the conflict. Though the scratching behavior cannot definitely be termed as an anxious behavior, evidence suggests that this is the case. An aggressor’s scratching decreases significantly after reconciliation. This suggests that reconciliation itself rather than a property of the conflict is the cause of reduction in scratching behavior. Though these results seem counter-intuitive, the anxiety of the aggressor appears to have a basis in the risks of ruining cooperative relationships with the opponent.[24]

Kin altruism and spite[edit]

In a study, a group of crab-eating macaques were given ownership of a food object. Unsurprisingly, adult females favored their own offspring by passively, yet preferentially, allowing them to feed on objects that they held. Interestingly, when juveniles were in possession of an object, mothers robbed them and acted aggressively at an increased rate towards their own offspring compared to other juveniles. These observations suggest that close proximity influences behavior in ownership, as a mother’s kin are closer to her on average. When given a nonfood object and two owners, one being a kin and one a non-kin, the rival will choose the older individual to attack regardless of kinship. Though the hypothesis remains that mother-juvenile relationships may facilitate social learning of ownership, the combined results clearly point to aggression towards the least threatening individual.[25]

A study was conducted in which food was given to 11 females. They were then given a choice to share the food with kin or non-kin. The kin altruism hypothesis suggests that the mothers would preferentially give food to their own offspring. Yet 8 out of the 11 females did not discriminate between kin and non-kin. The remaining 3 did, in fact, give more food to their kin. The results suggest that it was not kin selection, but instead spite that fueled feeding kin preferentially. This is due to the observation that food was given to kin for a significantly longer period of time than needed. The benefit to the mother is decreased due to less food availability for herself and the cost remains great for non-kin due to not receiving food. If these results are correct, crab-eating macaques are unique in the animal kingdom, as they appear not only to behave according to the kin selection theory but also act spiteful toward one another.[26]

Reproduction[edit]

After a gestation period of 162–193 days, the female gives birth to one infant. The infant's weight at birth is approximately 320 grams (11 oz).[12] Infants are born with black fur which will begin to turn to a yellow-green, grey-green, or reddish-brown shade (depending on the subspecies) after about three months of age.[16] This natal coat may indicate to others the status of the infant, and other group members treat infants with care and rush to their defense when distressed. Immigrant males will sometimes kill infants not their own, and high-ranking females sometimes kidnap the infants of lower-ranking females. These kidnappings usually result in the death of the infants, as the other female is usually not lactating. Young juveniles stay mainly with their mother and relatives. As male juveniles get older, they become more peripheral to the group. Here they play together, forming crucial bonds that may help them when they leave their natal group. Males that emigrate with a partner are more successful than those that leave alone. Young females, on the other hand, stay with the group and become incorporated into the matriline into which they were born.[27]

Male crab-eating macaques will groom females to increase the chance of mating. A female is more likely to engage in sexual activity with a male that has recently groomed her than with one that has not.[28]

Diet[edit]

A crab-eating macaque

Crab-eating macaques typically do not consume crabs; rather, they are opportunistic omnivores, eating a variety of animals and plants. Although fruits and seeds make up 60 - 90% of their diet, they also eat leaves, flowers, roots and bark.[12] They sometimes prey on vertebrates (including bird chicks, nesting female birds, lizards, frogs and fish), invertebrates and bird eggs. In Indonesia the species has become a proficient swimmer and diver for crabs and other crustaceans in mangrove swamps.[29]

This species exhibits particularly low tolerance for swallowing seeds. Despite their inability to digest seeds, many primates of similar size will swallow large seeds, up to 25 mm (0.98 in), and simply defecate them whole. The crab-eating macaque, on the other hand, spits seeds out if they are larger than 3–4 mm (0.12-0.16 in). This decision to spit seeds is thought to be adaptive; it avoids filling the monkey’s stomach with wasteful bulky seeds that cannot be used for energy.[30]

Although the species is ecologically well-adapted and poses no threat to population stability of prey species in its native range, in areas where the crab-eating macaque is not native, it can pose a substantial threat to biodiversity.[31] Some believe that the crab-eating macaque is responsible for the extinction of forest birds by threatening critical bird breeding areas [32] as well as eating the eggs and chicks of endangered forest birds.[18]

The crab-eating macaque can become a synanthrope, living off human resources. They are known to feed in cultivated fields on young dry rice, cassava leaves, rubber fruit, taro plants, coconuts, mangos and other crops, often causing significant losses to local farmers.[8] In villages, towns, and cities, they frequently take food from garbage cans and refuse piles.[8] The species can become unafraid of humans in these conditions, which can lead to macaques directly taking food from people, both passively and aggressively.

Tool use[edit]

Stone tool usage by crab-eating macaques in Laem Son National Park in Thailand.

In Thailand and Myanmar, crab-eating macaques use stone tools to open nuts, oysters and other bivalves, and various types of sea snails (nerites, muricids, trochids, etc.) along the Andaman sea coast and offshore islands.[13]

Another instance of tool use is washing and rubbing foods such as sweet potatoes, cassava roots, and papaya leaves before consumption. Crab-eating macaques either soak these foods in water or rub them through their hands as if to clean them. They also peel the sweet potatoes, using their incisors and canine teeth. Adolescents appear to learn these behaviors by observational learning of older individuals.[33]

Distribution and habitat[edit]

The crab-eating macaque lives in a wide variety of habitats, including primary lowland rainforests, disturbed and secondary rainforests, shrubland, and riverine and coastal forests of nipa palm and mangrove. They also easily adjust to human settlements; they are considered sacred at some Hindu temples and on some small islands,[9] but a pest around farms and villages. Typically, they prefer disturbed habitats and forest periphery. The native range of this species includes most of mainland Southeast Asia, from extreme Southeastern Bangladesh down through Malaysia, and the Maritime Southeast Asia islands of Sumatra, Java, and Borneo, offshore islands, the islands of the Philippines, and the Nicobar Islands in the Bay of Bengal. This monkey is a rare example of a terrestrial mammal that violates the Wallace Line.[18]

Introduced range[edit]

M. fascicularis is an introduced alien species in several locations, including Hong Kong, Taiwan, Irian Jaya, Papua New Guinea, New Britain, New Ireland, Pohnpei, Samoa, Tonga, Solomon Islands, Fiji, Nauru, Kiribati, Tuvalu, Vanuatu, Anggaur Island in Palau, and Mauritius. Where it is not a native species, particularly on island ecosystems whose species often evolved in isolation from large predators, M. fascicularis is a documented threat to many native species.[18] This has led the World Conservation Union (IUCN) to list M. fascicularis as one of the "100 worst invasive alien species".[34] Insofar as it is present as an invasive alien species on several islands, it has been labeled a "weed" species and is yet another significant ecological threat to those ecosystems and the species within them. However, M. fascicularis is not a threat to biodiversity in its native range.

The immuno-vaccine Porcine Zona Pellucida (PZP) which causes infertility in females is currently being trialed in Hong Kong to investigate its use as potential population control.[18]

Relationship with humans[edit]

A crab-eating macaque living in a human vicinity, with a stolen asthma inhaler.

Crab-eating macaques extensively overlap with humans across their range in Southeast Asia. Consequently, people and crab-eating macaques live together in many locations. Some of these areas are associated with religious sites and local customs, such as the temples of Bali in Indonesia, Thailand, and Cambodia, while other areas are characterized by conflict as a result of habitat loss and competition over food and space.[7] Humans and crab-eating macaques have shared environments since prehistoric times, and tend to both frequent forest and river edge habitats. Crab-eating macaques are occasionally used as a food source for some indigenous forest dwelling peoples. In Mauritius, they are captured and sold to the pharmaceutical industry, and in Angaur and Palau they are sold as pets.[18] Macaques feed on sugar cane and other crops, affecting agriculture and livelihoods, and can be aggressive towards humans. Macaques may carry potentially fatal human diseases, including Herpes B virus.[18]

In scientific research[edit]

M. fascicularis is also used extensively in medical experiments, in particular those connected with neuroscience and disease. Due to their close physiology, they can share infections with humans. Some cases of concern have been an isolated event of Reston ebolavirus found in a captive-bred population shipped to the US from the Philippines, which was later found to be a strain of Ebola that has no known pathological consequences in humans, unlike the African strains.[7] Furthermore, they are a known carrier of monkey B virus (Herpesvirus simiae), a virus which has produced disease in some lab workers working mainly with rhesus macaques (M. mulatta).[7] Nafovanny, the largest facility for the captive breeding of nonhuman primates in the world, houses 30,000 macaques .[citation needed] The crab-eating macaque is one of the types of monkeys that have been used as space test flight animals .[citation needed] It has been discovered recently that Plasmodium knowlesi, which causes malaria in M. fascicularis, can also infect humans. A few cases have been documented in humans, but for how long humans have been getting infections of this malarial strain is unknown. It is, therefore, not possible to assess if this is a newly emerging health threat, or if just newly discovered due to improved malarial detection techniques.[7] Given the long history of humans and macaques living together in SE Asia, it is likely the latter.

Conservation status[edit]

The crab-eating macaque has the third-largest range of any primate species, behind only humans and rhesus macaques. The IUCN Red List categorizes the species as Least Concern, and CITES lists them as Appendix II ("not necessarily threatened with extinction", in which trade must be controlled to avoid use incompatible with their survival). A recent review of their populations suggests a need for better monitoring of populations due to increased wild trade and rising levels of human-macaque conflict, which are reducing overall population levels despite the species being widely distributed.[35]

Each subspecies faces differing levels of threats, and there is too little information on some subspecies to assess their conditions. The M. f. umbrosa subspecies is likely of important biological significance and has been recommended as a candidate for protection in the Nicobar Islands, where its small, native population has been seriously fragmented,[36] and is listed as vulnerable on the IUCN Red List.[2] The Philippine long-tailed macaque (M.f. philippensis) is listed as near threatened, and M.f. condorensis is vulnerable.[2] All other subspecies are listed as data deficient and need further study;[2] although recent work is showing M.f aurea and M.f. karimondjawae need increased protection.[36] One concern for conservation is, in areas where M. fascicularis is not native, their populations need to be monitored and managed to reduce their impact on native flora and fauna.[36]

Genome[edit]

Genomic information
 NCBI genome ID  776
 Ploidy  diploid
 Genome size  2,946.84 Mb
 Number of chromosomes  21 pairs



The genome of the crab-eating macaque was sequenced.

See also[edit]

References[edit]

  1. ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 161–162. OCLC 62265494. ISBN 0-801-88221-4. 
  2. ^ a b c d e f Ong, P. & Richardson, M. (2008). Macaca fuscicularis. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 4 January 2009.
  3. ^ P H Napier, C P Groves (July 1983). "Simia fascicularis Raffles, 1821 (Mammalia, Primates): request for the suppression under the plenary powers of Simia aygula Linnaeus, 1758, a senior synonym. Z.N.(S.) 2399". Bulletin of Zoological Nomenclature 40 (2): 117–118. ISSN 0007-5167. Retrieved 19 November 2012. "Simia aygula is quite clearly the Crab-eating or Long-tailed Macaque, as Buffon opined as early as 1766." 
  4. ^ J. D. D. Smith (2001). "Supplement 1986-2000". Official List and Indexes of Names and Works in Zoology. International Trust for Zoological Nomenclature. p. 8. Retrieved 19 November 2012. "Suppressed under the plenary power for the purposes of the Principle of Priority, but not for those of the Principle of Homonymy" 
  5. ^ Wilson, D. E.; Reeder, D. M., eds. (2005). "Macaca fascicularis fascicularis". Mammal Species of the World (3rd ed.). Johns Hopkins University Press. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  6. ^ Linnaeus, Carl (1758). Systema naturæ. Regnum animale. (10 ed.). p. 27. Retrieved 19 November 2012. 
  7. ^ a b c d e f g Gumert, MD, Fuentes A, Jones-Engel, L. (2011). Monkeys on the Edge: Ecology and Management of Long-tailed Macaques and their Interface with Humans. Cambridge University Press. 
  8. ^ a b c Long, John (2003). Introduced Mammals of the World: Their History, Distribution, and Influence. Australia: CSIRO Publishing. p. 74. ISBN 0643067140. 
  9. ^ a b "Island of the Monkey God". Off the Fence. Retrieved 2013-09-25. 
  10. ^ a b van Noordwijk, Maria, and Carcel van Schaik (January 1999). "The Effects of Dominance Rank and Group Size on Female Lifetime Reproductive Success in Wild Long-tailed Macaques, Macaca fascicularis". Primates 40: 105–130. doi:10.1007/bf02557705. 
  11. ^ a b de Ruiter, Jan, and Eli Geffen (October 1998). "Relatedness of matrilines, dispersing males and social groups in long-tailed macaques (Macaca fascicularis)". Proceedings of the Royal Society B 285: 79–87. 
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  13. ^ a b Gumert, MD; Kluck, M.; Malaivijitnond, S. (2009). "The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand". American Journal of Primatology 71: 594–608. doi:10.1002/ajp.20694. 
  14. ^ Zimmer, Benjamin. "Makaku, macaco, macaque, macaca". Language Log. Retrieved 2013-09-26. 
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  16. ^ a b c d Cawthon Lang, Kristina. "Primate Factsheets: Long-tailed macaque (Macaca fascicularis) Taxonomy, Morphology, & Ecology". Primate Info Net. Retrieved 25 September 2013. 
  17. ^ Wilson, Don E.; Reeder, DeeAnn M., eds. (2005). "Macaca fascicularis". Mammal species of the world : a taxonomic and geographic reference (3rd ed. ed.). Baltimore: Johns Hopkins University Press. ISBN 978-0-8018-8221-0. 
  18. ^ a b c d e f g Carter, Steve. "Global Invasive Species Database". Macaca fascicularis (mammal). Invasive Species Specialist Group. Retrieved 26 September 2013. 
  19. ^ de Ruiter, Jan, Jan van Hooff, and Wolfgang Scheffrahn (June 1995). "Social and Genetic Aspects of Paternity in Wild Long-Tailed Macaques (Macaca fascicularis)". Behaviour 129: 203–224. 
  20. ^ Schaik, Carel, Maria Noordwijk, Rob Boer, and Isolde Tonkelaar (April 1983). "The effect of group size on time budgets and social behavior in wild long-tailed macaques". Behavioral Ecology and Sociobiology 13: 173–181. doi:10.1007/bf00299920. 
  21. ^ Cords, Marina (December 1992). "Post-conflict reunions and reconciliation in long-tailed macaques". Animal Behavior 44: 57–61. doi:10.1016/s0003-3472(05)80754-7. 
  22. ^ Hemelrijk, Charlotte (February 1994). "Support for being groomed in long-tailed macaques, Macaca fascicularis". Animal Behavior 48: 479–481. doi:10.1006/anbe.1994.1264. 
  23. ^ Aureli, Filippo (June 1992). "Post-conflict behaviour among wild long-tailed macaques, (Macaca fascicularis)". Behavioral Ecology and Sociobiology 31: 329–337. doi:10.1007/bf00177773. 
  24. ^ Das, Marjolijn, Zsuzsa Penke, and Jan van Hoof (1998). "Potconflict Affiliation and Stress-Related Behavior of Long-Tailed Macaque Aggressors". International Journal of Primatology 19: 53–71. 
  25. ^ Kummer, Hans and Marina Cords (December 1991). "Cues of ownership in long-tailed macaques, Macaca fascicularis". Animal Behavior 42: 529–549. doi:10.1016/s0003-3472(05)80238-6. 
  26. ^ Schaub, Hanspeter (1996). "Testing Kin Altruism in Long-Tailed Macaques (Macaca fascicularis) in a Food-sharing Experiment". International Journal of Primatology 17: 445–467. doi:10.1007/bf02736631. 
  27. ^ Cawthon Lang, Kristina. [<http://pin.primate.wisc.edu/factsheets/entry/long-tailed_macaque/behav> "Primate Factsheets: Long-tailed macaque (Macaca fascicularis) Behavior"]. Primate Info Net. Retrieved 26 September 2013. 
  28. ^ Gumert, Michael D. (December 2007). "Payment for sex in a macaque mating market". Animal Behavior 74 (6): 1655–1667. doi:10.1016/j.anbehav.2007.03.009. 
  29. ^ http://planet.uwc.ac.za/nisl/biodiversity/loe/page_289.htm
  30. ^ Corlett, R.T.; P.W. Lucas (1990). "Alternative Seed-Handling Strategies in Primates: Seed-Spitting by Long-Tailed Macaques (Macaca fascicularis)". Oecologia 82 (2): 166–171. doi:10.1007/bf00323531. Retrieved 25 September 2013. 
  31. ^ Richer, Nathan. "Wild Facts". Wild Fact #834 – The Perfect Gift – Crab-Eating Macaque. Retrieved 26 September 2013. 
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  33. ^ Wheatley, Bruce (June 1988). "Cultural behavior and extractive foraging in Macaca Fascicularis". Current Anthropology 29 (3): 516–519. doi:10.1086/203670. 
  34. ^ Lowe, S. "100 Of The World's Worst Invasive Species". Invasive Species Specialist Group. Retrieved 26 September 2013. 
  35. ^ Eudey, Ardith (2008). "The crab-eating macaque (Macaca fascicularis) widespread and rapidly declining.". Primate conservation 23: 129–132. doi:10.1896/052.023.0115. 
  36. ^ a b c Umapathy, G.; Singh M., Mohnot, S.M. (2003). "Status and Distribution of Macaca fascicularis umbrosa in the Nicobar Islands, India". International Journal of Primatology 24: 281–293. 


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